1. Introduction
The hydrodynamic behaviours and transport of microswimmers in a background flow are essentially relevant to planktonic active matter systems (Sengupta Reference Sengupta2023), aquatic ecology (Koehl & Cooper Reference Koehl and Cooper2015; Fuchs & Gerbi Reference Fuchs and Gerbi2016) and biological processes (Pedley, Hill & Kessler Reference Pedley, Hill and Kessler1988; Kuznetsov Reference Kuznetsov2011). Examples of such typical problems include the formation of microscale patches for motile phytoplankton in turbulence (Durham et al. Reference Durham, Climent, Barry, De Lillo, Boffetta, Cencini and Stocker2013), the process of giant whales in the ocean preying on zooplankton (Fenchel Reference Fenchel1988) and the red tide phenomenon caused by microbial communities (Zohdi & Abbaspour Reference Zohdi and Abbaspour2019). In lakes and oceans, these microorganisms must adapt their motility to the surrounding flows to find food, escape predators, produce oxygen, optimize light uptake and mate.
An individual motile planktonic microorganism in a background flow often swims in a preferred direction due to a stabilizing torque that can arise from an asymmetry in shape or body density or the ability to sense the direction of gravity, and the viscous torque exerted on cells by fluid shear (e.g. the velocity gradient of the background flow field) (Pedley & Kessler Reference Pedley and Kessler1990, Reference Pedley and Kessler1992). These two torques act to overturn cells competitively or collaboratively. As the number of microorganisms increases, the microswimmers are known to exhibit intriguing collective behaviours, including small-scale clustering, vertical trapping and enhanced or suppressed transport compared with passive spheres, which has attracted considerable attention recently (Ruiz, Macìas & Peters Reference Ruiz, Macías and Peters2004; De Lillo et al. Reference De Lillo, Cencini, Durham, Barry, Stocker, Climent and Boffetta2014; Gustavsson et al. Reference Gustavsson, Berglund, Jonsson and Mehlig2016; Borgnino et al. Reference Borgnino, Boffetta, De Lillo and Cencini2018; Cencini et al. Reference Cencini, Boffetta, Borgnino and De Lillo2019; Marchioli et al. Reference Marchioli, Bhatia, Sardina, Brandt and Soldati2019; Qiu, Marchioli & Zhao Reference Qiu, Marchioli and Zhao2022; Yang et al. Reference Yang, Jiang, Picano and Zhu2024). Torney & Neufeld (Reference Torney and Neufeld2007) found that the swimming velocity of prolate swimmers has a threshold, dependent on the aspect ratio, for no-barrier transport. Using the classical Vicsek flocking model to simulate microswimmers in turbulent flow, a recent study suggests that realistic self-propelling dynamics must be considered when assessing the stability of collective behaviour (Khurana & Ouellette Reference Khurana and Ouellette2013). Baggaley (Reference Baggaley2016) found that the critical balance between alignment with neighbouring particles and anticipation of their motion stabilizes the flock in turbulent flows. Durham et al. (Reference Durham, Climent, Barry, De Lillo, Boffetta, Cencini and Stocker2013) reported that Heterosigma akashiwo forms striking patches within individual vortices experimentally and proved with a mathematical model that this patchiness originated from the coupling between motility and shear. Fluid turbulence plays a significant role in the collective dynamics of swimming anisotropic particles in their accumulation and reorientation. Zhan et al. (Reference Zhan, Sardina, Lushi and Brandt2014) studied how microswimmers’ velocity and aspect ratio affect particle clustering in isotropic turbulence. They found that a three-dimensional turbulent flow significantly reduces the clustering and patchiness that widely exists in laminar flows, linear shear and vortex flows. It was concluded that this is due to the complex flow topology of turbulent flows. Additionally, elongated microorganisms show some level of clustering in the case of swimmers without any preferential alignment, whereas spherical swimmers remain uniformly distributed. Subsequently, Pujara, Koehl & Variano (Reference Pujara, Koehl and Variano2018) investigated the effects of swimmers’ shapes (sphere, rod, disc and triaxial ellipsoid) on the accumulation in isotropic turbulence. It was found that the particle enstrophy (variance of particle angular velocity) is largest for triaxial ellipsoids swimming at a velocity smaller than the Kolmogorov velocity scale; the swimmers tend to accumulate at regions with lower fluid vorticity compared with passive particles; and the average alignment of particles with the directions of the velocity gradient tensor is altered by swimming leading to an equal distribution of rotation about different particle axes. Recently, Zhang, Qiu & Zhao (Reference Zhang, Qiu and Zhao2022) studied the collective behaviours of microswimmers in vertical turbulent channel flows, finding that the swimmers distribute non-uniformly in the wall-normal direction and with preferential orientation in the mean flow direction. Swimmers with a predominant swimming velocity tend to accumulate near the solid walls, regardless of the flow direction. They explained the wall-normal transport process, including particle–fluid slip velocity and local clustering.
The above efforts usually assume that the effect of finite microswimmer size is negligible (a point-particle model), and the interaction between the microswimmers as well as the reversed effect of motile microswimmers on the turbulent flows have yet to be considered. These simplifications are justified when the swimmer dimensions are far smaller than the Kolmogorov scale of the flow, and the volume fractions of the swimmers are low enough to satisfy the assumption of dilute suspension in the bulk of the flow. In the open oceans and shallow coastal waters, the turbulent kinetic energy dissipation rate ε at 20 °C respectively achieves ranges of 10−11–10−6 W kg−1 (Thorpe Reference Thorpe2005) and 10−9–10−4 W kg−1 (Bluteau, Jones & Ivey Reference Bluteau, Jones and Ivey2016; Masunaga et al. Reference Masunaga, Fringer, Yamazaki and Amakasu2016), corresponding to the Kolmogorov scales of 1.0–17.8 mm and 300–500 μm. Hence, for many phytoplankton with sizes smaller than 5 μm, the point-particle model may be valid for simulations. However, the scales of phytoplankton usually fall into the range O(0.1–1000 μm) (Guasto, Rusconi & Stocker Reference Guasto, Rusconi and Stocker2012; Sengupta Reference Sengupta2023), and some of the scales (e.g. the green alga Volvox is about 500 μm in length) are around the Kolmogorov scale. Moreover, as an important component of plankton, zooplankton (e.g. ostracods, mysids, polychaetes, decapods, monstrilloid copepods and hydrozoans) is usually larger than phytoplankton, with their sizes reaching 0.7–5 mm (Kiørboe, Jiang & Colin Reference Kiørboe, Jiang and Colin2010; Tanaka et al. Reference Tanaka, Genin, Endo, Ivey and Yamazaki2020). This scale is comparable to or even greater than the Kolmogorov scale in the oceans, with the scale ratio (zooplankton length to Kolmogorov scale) being in the range of approximately 0.04–17. Consequently, the point-particle model is no longer applicable when considering the swimming of large phytoplankton or zooplankton in turbulent flows here. This requires the swimmers’ geometry to be resolved. Apart from the scale issues, it is noteworthy that the available investigations have primarily focused on the swimming falling into the limit of the Stokes flow regime (neglecting the effect of swimming fluid inertia). Actually, many medium- or giant-sized plankton swimming achieves a finite Reynolds number (Re) in the range O(1–10) in escape or from predators (Childress Reference Childress and Childress1981; Beckett Reference Beckett1986; Guasto et al. Reference Guasto, Rusconi and Stocker2012). For example, in the natural environment, Re is in the range 0.1–1 for freely swimming zooplankton Daphnis magna (Wickramarathna, Noss & Lorke Reference Wickramarathna, Noss and Lorke2014) and 0.2–2 for paramecia depending on swimming or escaping mode (Ishikawa & Hota Reference Ishikawa and Hota2006), and the millimetre-sized copepods and Pleurobrachia commonly achieve Re of 10 (Kiørboe et al. Reference Kiørboe, Jiang and Colin2010). This swimming is beyond the Stokes flow regime, and the fluid inertia can remarkably affect the microswimmers’ motion, either enhancing or hindering their speed (Wang & Ardekani Reference Wang and Ardekani2012; Khair & Chisholm Reference Khair and Chisholm2014; Ouyang, Lin & Ku Reference Ouyang, Lin and Ku2018; More & Ardekani Reference More and Ardekani2020), modifying the contact time between a swimmer with solid boundaries and another swimmer (Li & Ardekani Reference Li and Ardekani2014; Ouyang, Lin & Ku Reference Ouyang, Lin and Ku2019) and responsible for the stability of flocks (Choudhary, Venkataraman & Ray Reference Choudhary, Venkataraman and Ray2015; Gupta et al. Reference Gupta, Roy, Saha and Ray2020). This indicates that mobile microorganisms exhibit collective behaviours in turbulent fields distinctly different from those of passive particles (Zhao et al. Reference Zhao, Challabotla, Andersson and Variano2015; Zhao & Andersson Reference Zhao and Andersson2016).
Based on the above situations, one may raise several new questions: (i) What collective behaviour do many motile microswimmers exhibit in turbulence when their scale is comparable to or greater than the Kolmogorov scale? (ii) How can these motile microswimmers modify turbulence in reverse? (iii) How does the fluid inertia of these motile microswimmers affect their collective behaviour in turbulence?
In this paper, we employ a classical spheroidal ‘squirmer’ model to investigate the collective behaviours of motile microswimmers in a turbulent channel flow. This microswimming model, established by Lighthill (Reference Lighthill1952) and extended by Blake (Reference Blake1971), is widely adopted to mimic the self-propulsion of a swimmer with a dense array of cilia on its surface. In contrast to the point-particle model, a squirmer is required to resolve the interface of the microswimmers fully, and it can induce the flow field around both puller- and pusher-type microswimmers by adjusting a swimming parameter (Ishikawa & Hota Reference Ishikawa and Hota2006; Chisholm et al. Reference Chisholm, Legendre, Lauga and Khair2016). It has successfully simulated many microswimming scenarios, including microorganisms’ nutrient uptake (Magar, Goto & Pedley Reference Magar, Goto and Pedley2003; Magar & Pedley Reference Magar and Pedley2005), their swimming in a viscoelastic fluid (Ouyang et al. Reference Ouyang, Lin, Lin, Phan-Thien and Zhu2023), hydrodynamic interactions with boundaries (Ishimoto & Gaffney Reference Ishimoto and Gaffney2013), two-body hydrodynamic interactions (Ishikawa, Simmonds & Pedley Reference Ishikawa, Simmonds and Pedley2006; Götze & Gompper Reference Götze and Gompper2010; Navarro & Pagonabarraga Reference Navarro and Pagonabarraga2010; Li, Ostace & Ardekani Reference Li, Ostace and Ardekani2016) and collective swimming dynamics (Ishikawa & Pedley Reference Ishikawa and Pedley2008; Ishikawa, Locsei & Pedley Reference Ishikawa, Locsei and Pedley2008; Zöttl & Stark Reference Zöttl and Stark2014; Ishikawa, Brumley & Pedley Reference Ishikawa, Brumley and Pedley2021; Zhang et al. Reference Zhang, Yu, Ouyang, Xia and Lin2024). Turbulent channel flow exemplifies the universality of the scenario in relevant environments, characterized by (i) varying turbulence scales across the channel, from the boundary to the centre, (ii) the presence of unidirectional shear flow and (iii) the influence of wall constraints. These characteristics are commonly found in natural and industrial settings where microswimmers operate. Examples include the movement of planktonic microorganisms in sewage pipelines (Muñoz & Guieysse Reference Muñoz and Guieysse2006), the shear flow experienced by mobile microorganisms on the seabed (Wheeler et al. Reference Wheeler, Secchi, Rusconi and Stocker2019) and the wall shear affecting yeast clusters during food fermentation processes (Sharma, Malhotra & Rathore Reference Sharma, Malhotra and Rathore2011). This combination of universal applicability and practical significance makes turbulent channel flow an ideal model when exploring the dynamics of microswimmers.
This paper utilizes a direct-forcing fictitious domain (DF-FD) method to investigate the hydrodynamic behaviours of a turbulent channel flow filled with motile microswimmers. We aim to elucidate how the microswimmers’ volume fraction, self-propelled mode and swimming Reynolds number affect their collective behaviours encompassing distribution, orientation and rotation. Additionally, we aim to illustrate how the presence of microswimmers modifies turbulence behaviours, including fluid velocity, stress and vortex structure. The remainder of this paper is organized as follows. Section 2 provides a brief overview of squirmer dynamics within the DF-FD framework. Following this, in § 3, we carefully validate the single-phase turbulent channel flow against the available data and establish the calculation parameters. Section 4 presents and discusses the results, including the fluid velocity and particle volume fraction profiles, flow drag and turbulent behaviours, squirmer orientation and rotation and vortex structure. Finally, in § 5, we offer some concluding remarks.
2. Numerical strategy
2.1. The DF-FD method
In the present study, we employ a DF-FD method proposed by Yu & Shao (Reference Yu and Shao2007) to conduct interface-resolved direct numerical simulation of turbulent channel flows. The basic strategy of this method is that the interior of the body (e.g. solid squirmer) is filled with a fictitious fluid, and a pseudo-body force is considered over the body’s inner domain to enforce the fictitious fluid to satisfy the rigid-body motion constraint hence embodying hydrodynamic interactions between the body and the fluid (Glowinski et al. Reference Glowinski, Pan, Hesla and Joseph1999). Briefly, we demonstrate in the following a non-dimensional FD formulation for the motion of an individual particle in an incompressible fluid. Let P 0 denote the solid domain and Ω 0 the entire domain including the interior and exterior of the solid body. We adopt the following scales for non-dimensionalization: H for length, U 0 for velocity, H/U 0 for time and ρ f U 0 2 for the pseudo-body force with ρ f being the fluid density. Then, the fluid continuity and momentum equations can be expressed as
where u and p are the fluid velocity and pressure, respectively;
$\boldsymbol\lambda$
is the vectorial Lagrange multiplier (pseudo-body force); and the Reynolds number is defined by Re = ρ
f
U
0
H/
$\mu$
. The fluids in the squirmer domain satisfy the rigid-body motion constraints, and the motion of the squirmer is governed by Newton’s second law:
where r is the position vector concerning the squirmer’s centreof mass; U and
ω
s
are its translational and angular velocities; u
s
denotes the velocity distribution for the squirmer dynamics which we will introduce later; ρ
r
is the squirmer–
fluid density ratio, ρ
r
= ρ
s
/ρ
f
, here ρ
r
= 1;
$V_{p}{}^{\!\ast}$
is the dimensionless volume defined by
$V_{p}{}^{\!\ast}$
= Vp
/H
3 with Vp
being the squirmer’s volume; and J
* is the dimensionless moment of inertia with J
* = J/ρ
s
H
5. We decouple equations (2.1)–(2.5) into the fluid and particle sub-problems using a fractional-step time method. For more details on the discrete schemes, the reader
is referred to the work of Yu & Shao (Reference Yu and Shao2007). In this study, we discretize the convection term using the MUSCL (high-precision monotone upstream-centred scheme for conservation laws) scheme, and the fourth-order Runge–Kutta method is employed for time integration.
2.2. Squirmer dynamics
The squirmer model introduces a progressive waving envelope to mimic both radial and angular oscillations at the boundary of a microswimmer with arrays of cilia like Volvox (Pedley, Brumley & Goldstein Reference Pedley, Brumley and Goldstein2016). A reduced-order squirmer, considering the pure steady tangential motion on its surface, has been extensively adopted to investigate the microswimming mechanisms under various background flows. This reduced-order model gives a tangential boundary condition at a spherical body’s surface (in the framework of reference moving with the body) for self-propelling, and it reads
where θ is the angle concerning the swimming direction and B 1 and B 2 are the first two squirming modes (parameters). Vector e θ is the unit vector along the polar direction. The squirmer can be categorized into a puller (β> 0; e.g. Chlamydomonas), pusher (β< 0; e.g. E. coli) and neutral squirmer (β = 0), based on the values of β = B 2/B 1 (B 1 > 0) (Ishikawa & Pedley Reference Ishikawa and Pedley2008). In the Stokes flow regime, the steady swimming speed of a squirmer through a Newtonian fluid is U 0 = 2B 1/3 (Lighthill Reference Lighthill1952). In the framework of the present DF-FD method, a solenoidal volumetric velocity u s is derived (using the boundary condition (2.6)) and exerted at the whole spherical solid domain (inside) to realize the self-propelling of the body (Li et al. Reference Li, Ostace and Ardekani2016; Lin & Gao Reference Lin and Gao2019; More & Ardekani Reference More and Ardekani2020; Ouyang et al. Reference Ouyang, Lin, Yu, Lin and Phan-Thien2022), and it reads
where a is the radius of the spherical squirmer; r is the distance from the squirmer’s centre; e r is the unit vector along the radial direction; and m is an arbitrary positive integer, where , usually, m = 5 is adopted. The motion of a spheroidal squirmer is governed by (2.4) and (2.5). The reader is referred to our previous work (Lin & Gao Reference Lin and Gao2019; Ouyang et al. Reference Ouyang, Lin, Yu, Lin and Phan-Thien2022) for the details of the squirmer dynamics in the framework of the DF-FD method.
2.3. Collision model
For many squirmers in a turbulent channel flow, the squirmers will collide with each other or the wall. In addition to hydrodynamic interactions, we implement a soft-sphere collision model to resolve the short-ranged collisions. It calculates a repulsive force between the ith and jth objects (squirmers and walls) that are close to contact (Lin & Gao Reference Lin and Gao2019):
where the constant κ 0 is typically chosen as 103–104 of particle mass; d is the particle interdistance; dc is a cutoff distance to define the ‘contact’ zone, which is dependent on mesh size Δx (typically dc = Δx–2Δx); and n is a unit vector pointing in the direction where contact occurs.
Schematic of a turbulent channel flow filled with finite-size microorganisms. The mean flow is along the x-axis direction.

3. Validation and simulation set-up
A schematic diagram of the channel flow is shown in figure 1. The no-slip velocity boundary condition is imposed on the channel walls (the y axis), and the periodic boundary condition is imposed in the streamwise (x) and spanwise (z) directions. As shown in figure 1, we employ a computational domain with L× R × W = [0, 8H] × [−H, H] × [0, 4H], where H is the half-channel width. We perform direct numerical simulations for all the cases, and the channel flow is driven by the mean pressure gradients along the streamwise direction. Here, we adopt H as the characteristic length and the friction velocity
$u_{\tau }=\sqrt{\tau _{w}/\rho _{f}}$
as the characteristic velocity. Note that τ
w
is the mean shear stress on the walls and ρ
f
is the fluid density. Hence, the friction Reynolds number is defined by Re
τ
= u
τ
H/ν, with ν being the fluid kinematic velocity. The corresponding bulk-flow Reynolds number is around Re
b
= u
max
H/ν ≈ 3320, where u
max
denotes the maximum flow velocity. After careful grid tests and referencing previous work using the present code (Yu et al. Reference Yu, Lin, Shao and Wang2016; Zhu, Yu & Shao Reference Zhu, Yu and Shao2018; Yu et al. Reference Yu, Xia, Guo and Lin2021), a computational mesh of 512 × 128 × 256 is employed for validating the single-phase turbulent channel flow. As shown in figure 2, our statistics of the carrier fluid, including the mean-velocity profiles and root-mean-square (RMS) velocity fluctuations, show good agreement with the results of Kim, Moin & Moser (Reference Kim, Moin and Moser1987) at Re
τ
= 180. Note that the half-channel width in figure 2 is non-dimensionalized by the viscous length scale ν/u
τ
, and denoted by a superscript plus sign. Note that we adopted a uniform mesh in the streamwise, spanwise and wall-normal directions, respectively, with a grid resolution of Δx
+ = Δz
+ = Δy
+ = 0.356.
Statistics of the carrier fluid. (a) Mean-velocity profiles and (b) RMS velocity fluctuations.

In this paper, we simulate spherical squirmers subjected to turbulent channel flows. To characterize the self-propelling behaviour of the squirmer, a swimming Reynolds number is defined as Re
s
= ρ
f
U
0
a/
$\mu$
, where the characteristic velocity is adopted as U
0 = 2B
1/3 (see (2.6)). In this study, the friction Reynolds number is set to Re
τ
= 180. Typical swimming Reynolds number Re
s
= 5 and 10 are adopted to characterize the swimming in a finite-flow regime if not otherwise specified. Given these settings, one can estimate a typical real-world scenario (Kiørboe Reference Kiørboe2011; Svetlichny at al. Reference Svetlichny, Larsen and Kiørboe2020; Tanaka et al. Reference Tanaka, Genin, Endo, Ivey and Yamazaki2020): microorganisms with a characteristic length of 1 mm and swimming speeds of 10 mm s−1 (Re
s
= 5) or 20 mm s−1 (Re
s
= 10) navigate within a turbulent channel that has a height of 8 mm and a maximum velocity of 83 mm s−1 at the channel centre. The non-dimensionalized Kolmogorov length η for this turbulent channel flow is calculated by
\begin{equation} \eta ^{*}=\frac{\eta }{H}=\frac{1}{H}\left(\frac{v^{3}}{\varepsilon }\right)^{1/4}=\frac{1}{H}\Bigg(\frac{v^{3}}{\varepsilon ^{*}\left({u_{\tau }}^{4}/v\right)}\Bigg)^{1/4} ,\end{equation}
where ε is the turbulence dissipation rate and ε
∗ is the dimensionless turbulence dissipation rate normalized with u
τ
4/v. Based on the available results (Hoyas & Jiménez Reference Hoyas and Jiménez2008; Yu et al. Reference Yu, Xia, Guo and Lin2021) and our calculation at Re
τ
= 180, the non-dimensionalized Kolmogorov length
$\eta ^{*}$
from the wall to the channel centre is estimated at the range of 0.0086–0.021. The non-dimensionalized viscous length is defined as l
v
* = l
v
/H = v/(u
τ
H) = 0.0056. Given this parameter setting, the scale ratio of Kolmogorov length (a/
$\eta$
) from the channel centre to the wall ranges between 5.95 and 14.5. Additionally, the scale ratio of the viscous length a/l
v
equals 22.5, where a is the squirmer’s radius. Furthermore, two velocity scale ratios involve friction and the mean maximum flow velocities. The ratios U
0/u
τ
are 0.22 (Re
s
= 5) and 0.44 (Re
s
= 10), and U
0/u
max
are 0.012 (Re
s
= 5) and 0.024 (Re
s
= 10), respectively. The calculation parameters for the typical cases are summarized in table 1. Note that a mesh size of 16Δx across the squirmer’s diameter and a time step of Δt = 5 × 10−4, which can afford the convergent results for the squirming problem (Lin & Gao Reference Lin and Gao2019), are adopted for the present study. Nevertheless, we also simulated the pusher (β = −5, Re
s = 5) example with a particle diameter resolved by 32 grids. Compared with results obtained using a grid resolution of 16, the differences were minimal, with relative errors occurring primarily at the peak of particle distribution and remaining below 1.5 % (see figure 4
a in the next section). This suggests that a grid resolution of 16 is sufficient to achieve convergent results.
Parameter settings for the typical cases studied. Here φ 0 denotes the average particle volume fraction through the entire channel and Np represents the particle number.

4. Results and discussion
4.1. Fluid velocity and particle volume fraction profiles
Figure 3 presents the mean fluid velocity profiles of the turbulent channel flows. Our findings support previous conclusions that suspended passive spheres decrease velocity across the entire profile even though the blocking ratio (the particle diameter to the channel width) here is larger than that in the previous study (Zhu et al. Reference Zhu, Yu and Shao2018). At a high volume fraction (𝜑0 = 12.7 %; see figure 3 a), the squirmers exhibit a similar feature to passive spheres in decreasing the velocity profile. However, the extent of this decrease varies among different types of squirmers. These differences may be relevant to the distribution of particles near the wall, as shown in figure 4(a). The peak of the particle volume fraction near the wall indicates the generation of a particle layer, resulting in drag enhancement (Picano, Breugem & Brandt Reference Picano, Breugem and Brandt2015; Costa et al. Reference Costa, Picano, Brandt and Breugem2016). Notably, a higher peak corresponds to a higher density of the particle layer, thus creating slower velocity profiles. Note that these peaks appear at the exact location (y +≈ 42) where there is a sudden weakening tendency in the velocity increase. For Chlamydomonas microalgae swimming in still fluids and near a wall, there is also a notable peak in their time-averaged concentration profile, approximately equal to the sum of the cell radius and the flagellar length (Contino et al. Reference Contino, Lushi, Tuval, Kantsler and Polin2015; Ostapenko et al. Reference Ostapenko, Schwarzendahl, Böddeker, Kreis, Cammann, Mazza and Bäumchen2018). In contrast, the present peak’s distance from the wall indicates the thickness of the particle layer, which is about 1.9 times the squirmer radius from the wall to the body centre. This thickness remains consistent regardless of the self-propulsion modes or the particle volume fraction.
Fluid mean velocity profiles of squirmer- and particle-laden turbulent channel flows at a/H = 0.125, Re τ = 180 for (a) 𝜑0 = 12.7 % and (b) 𝜑0 = 2.54 %.

Profiles of the particle volume fraction at a/H = 0.125, Re τ = 180 for (a) 𝜑0 = 12.7 % and (b) 𝜑0 = 2.54 %. The vertical axis is dimensionless by the volume fraction of the squirmers. Pusher1 indicates a grid resolution of 32 across its diameter.

Regarding the profiles of the squirmer volume fraction at Re s = 5, the peak for the pushers is the largest, followed by that for the neutral squirmers and subsequently for pullers. Consequently, the higher peak results in a slower velocity profile within squirmers. This distribution pattern near the wall (see the peaks) is different from the collective behaviours of multiple squirmers between two walls (without external channel flows), in which pushers and pullers have stronger tendencies to accumulate near the wall compared with neutral squirmers (Li & Ardekani Reference Li and Ardekani2014). The disparity between the two lies in the dominance of distinct mechanisms. In a quiescent fluid, the collective behaviour of particles is dictated by their individual motion and hydrodynamic interactions. Hence, the pullers and pushers tend to accumulate near the wall due to the presence of stresslet (strongest far-field interaction), in contrast to the neutral squirmers lacking this characteristic (Spagnolie & Lauga Reference Spagnolie and Lauga2012). However, in this context, the distribution of squirmers is largely determined by the turbulent channel flows, leading to the intense collisions of spherical particles in the near-wall region (Zhu et al. Reference Zhu, Yu and Shao2018). Compared with passive spheres, squirmers generally result in a further weakening of fluid velocity. However, a notable exception arises with pullers, which instigate a slight augmentation in fluid velocity within the bulk-flow region at Re s = 5 (y + ≥ 75; see figure 3 a). On further increasing Re s to 10, the effect of pullers does not exhibit amplification, as evidenced by a peak in figure 4(a) that is higher than that observed at Re s = 5.
At a low volume fraction (𝜑0 = 2.54 %), as shown in figure 3(b), comparable yet diminished effects are discerned in the impeding of velocity profiles by squirmers in contrast to those noted at a high volume fraction. Notably, in comparison with passive spheres, a low volume fraction of pullers (Res = 5) results in a slightly enhanced velocity profile across the channel. This observation is consistent with the finding that the peak of the passive particle distribution exceeds that of the pullers (see figure 4 b), contrasting with the trend observed at high volume fraction. The reduction in particle volume fraction (from 𝜑0 = 12.7 % to 2.54 %) leads to a transition in the distribution profile of particles from bimodal to unimodal. This change can be attributed to variations in hydrodynamic interactions. As shown in figure 4(a), the primary peak of particle distribution, from the vicinity of the wall to the centre of the channel, is associated with particle–wall interaction. In contrast, the secondary peak corresponds to particle–particle interaction. Consequently, as particle volume fraction decreases, the strength of particle–particle interaction diminishes significantly, resulting in the disappearance of the secondary peak, as depicted in figure 4(b).
Compared with the point-particle model demonstrating microswimmers (it is valid for particle dimensions far smaller than the Kolmogorov scale of the flow), our finite-sized squirmers are more inclined towards the wall. Zhang et al. (Reference Zhang, Qiu and Zhao2022) suggested that increasing swimming speed (related to the present Re s ) results in a slight migration towards the bulk-flow region for spherical microswimmers, and there is no noticeable peak (point-particle model). In contrast, the increase of Re s from 5to 10 strengthens the aggregate behaviours of the pullers in the near-wall region, leading to the hindrance of the velocity profile (see figures 3 and 4). This discrepancy may be attributed to the near-field flow fields induced by the microswimmers. Li & Ardekani (Reference Li and Ardekani2014) concluded that an individual puller is more easily entrapped near the wall at a higher Res due to the presence of stresslet, qualitatively having interpreted our results. For the point-particle model, the microscopic hydrodynamic interactions between particles and the wall have not been fully resolved, and different types of microswimmers are not distinguished. Hence, the collective behaviours are primarily affected by the external turbulent flows.
Profiles of the fluid viscous stress τ fV , the fluid Reynolds stress τ fR , the particle Reynolds stress τ pR and the particle inner stress τ pI at φ 0 = 12.7 % (lines) and 2.54 % (circles). (a) Spheres. (b) Neutral squirmers (β = 0, Re s = 5). (c) Pushers (β = −5, Re s = 5). (d) Pullers (β = 5, Re s = 5 and 10, φ 0 = 12.7 %). The stresses τ + are normalized by ρ f u τ 2.

4.2. Flow drag and turbulent behaviours
To analyse the contributions of flow drag, we decouple the total stress of the channel flows filled with squirmers based on the mean momentum equation (Picano et al. Reference Picano, Breugem and Brandt2015; Zhu et al. Reference Zhu, Yu and Shao2018), and it reads
\begin{equation} \underset{\tau _{T}}{\underbrace{\tau _{w}\left(1-\frac{y}{H}\right)}}=\underset{\tau _{fV}}{\underbrace{\varphi _{f}\mu \frac{{\textrm d}\langle u_{xf}\rangle }{{\textrm d}y}}}+\underset{\tau _{pI}}{\underbrace{\varphi _{s}\langle \sigma _{p}\rangle _{xy}}}+\underset{\tau _{fR}}{\underbrace{\varphi _{f}\rho _{f}\langle -u_{xf}^{\prime}u_{yf}^{\prime}\rangle }}+\underset{\tau _{pR}}{\underbrace{\varphi _{s}\rho _{s}\langle -u_{xp}^{\prime}u_{yp}^{\prime}\rangle }} ,\end{equation}
where u
x
and u
y
are, respectively, the velocity components in the x and y directions; the subscripts f and p represent the fluid phase and solid phase, respectively; σ
p
denotes the solid inner stress; and
$\varphi _{f}$
and
$\varphi _{s}$
are the local volume fractions of fluid and solid, respectively. Note that the angle brackets denote the phase averaging of fluid or solid. The five terms in (4.1) denote the total stress τ
T
, the fluid viscous shear stress τ
fV
, the particle inner stress τ
pI
, the fluid Reynolds stress τ
fR
and the particle Reynolds stress τ
pR
. Figure 5 presents the contributions of the total stress along the normal direction of the wall. The fluid viscous shear stress τ
fV
decreases monotonically from the wall to the channel centre, mirroring the analogous trend observed in viscous dissipation. The main finding in this section is that the presence of different self-propelling modes can significantly affect the particle inner stress τ
pI
in the near-wall regions differently, particularly pronounced in scenarios characterized by high particle volume fraction (𝜑0 = 12.7 %). At 𝜑0 = 12.7 %, the pushers contribute to the highest peak of τ
pI
among the squirmers, followed by that of the neutral squirmers and subsequently that of pullers. This pattern may be attributed to the joint impacts of particles’ accumulation behaviours (volume fraction) and the self-propelling features. Predominantly, the elevated peak of τ
pI
correlates with the heightened peak of the particle volume fraction in the near-wall region (see figure 4). Additionally, a faster squirmer results in a larger τ
pI
(the steady swimming speeds at Res
= 5 in descending order are a pusher (β = −5), neutral squirmer (β = 0) and puller (β = 5) (Chisholm et al. Reference Chisholm, Legendre, Lauga and Khair2016); for instance, even though the particle volume fraction of pushers in the bulk flow region is smaller than that of passive spheres, pullers and neutral squirmers, τ
pI
of the pusher case is still larger than that of other cases). The particle Reynolds stress τ
pR
in flows containing squirmers exhibits a slight increase compared with those involving passive spheres. This can be attributed to implementing the self-propelling velocity boundary condition as (2.7). Significantly, the increase in τ
pI
is primarily at the expense of diminishing the fluid Reynolds stress τ
fR
. This pattern is also replicated in figure 5(d), and the increase in Res
(from 5to 10) results in the enhancement of τ
pI
accompanied by a reduction in τ
fR
.
The conclusion outlined above broadly applies to the low particle volume fraction. At 𝜑0 = 2.54 %, these stresses are further integrated into figure 6 for a more comprehensive comparison. The variation in fluid viscous stress induced by different squirmers is minimal. In contrast, the particle Reynolds stress τ pR at a specified Reynolds number (Re s = 5) demonstrates a positive correlation with the squirmers’ steady speeds (Chisholm et al. Reference Chisholm, Legendre, Lauga and Khair2016). However, two notable findings emerge at a low concentration, distinct from those observed at a high concentration. Firstly, the peak of particle inner stress τ pI for the puller case is smaller than that of passive spheres at 𝜑0 = 2.54 % (Re s = 5), resulting in a greater fluid Reynolds stress τ fR . This observation could be a key factor contributing to the higher flow profile velocity observed for pullers compared with passive spheres (see figure 3 b). Secondly, at 𝜑0 = 2.54 %, an increase in the swimming Reynolds number of the pullers (from 5to 10) weakens particle inner stress (enhancing fluid Reynolds stress). Interestingly, this variation with Re s does not translate into an increase in the velocity profile, in contrast to the findings at 𝜑0 = 12.7 %. This inconsistency may be attributed to the fact that the increase in Res enhances the concentration of pullers near the wall more significantly at 𝜑0 = 12.7 % than at 2.54 % (see figure 4). Consequently, when increasing Re s from 5to 10, the puller case with 𝜑0 = 12.7 % causes a more significant hindrance in the velocity profile, while those with 𝜑0 = 2.54 % do not.
Comparison of the fluid stress at φ 0 = 2.54 %. (a) Viscous stress τ fV and particle Reynolds stress τ pR . (b) Fluid Reynolds stress τ fR and the particle inner stress τ pI .

To quantify the effects of individual stresses on flow drag, the relationship between the bulk velocity and stresses derived from the mean momentum equation (4.1) reads
Contributions of the individual stresses to the flow drag for different squirmers (φ 01 = 12.7 %, φ 02 = 2.54 %).

The bulk velocity of the entire particle suspension flows can be calculated by integrating (4.2) twice:
\begin{align} u_{b}&=\frac{1}{H}\int _{0}^{H}(\varphi _{f}\langle u_{f}\rangle +\varphi _{s}\langle u_{s}\rangle ){\textrm{d}}y \nonumber\\ &=\frac{1}{H}\int _{0}^{H}\varphi _{f}\int _{0}^{y}\frac{1}{\mu \varphi _{f}}(\tau _{T}-\tau _{fR}-\tau _{pI}-\tau _{pR})\;{\textrm{d}}\xi \textrm{dy}+\frac{1}{H}\int _{0}^{H}(\varphi _{s}\left\langle u_{s}\right\rangle ){\textrm{d}}y. \end{align}
Equation (4.3) can be normalized by the characteristic length H and the characteristic velocity
$u_{\tau }$
, and we follow the definition of Zhu et al. (Reference Zhu, Yu and Shao2018) with the dimensionless terms of the total stress, the fluid Reynolds stress, the particle inner stress, the particle Reynolds stress and the particle mean velocity as C
T
, C
fR
, C
pI
, C
pR
and C
up
, respectively. The dimensionless values are summarized in
table 2. Note that the total stress term in the single-phase scenario is related to the friction Reynolds number, expressed as C
T
= Reτ
/3 (Yu et al. Reference Yu, Lin, Shao and Wang2017). At 𝜑0 = 12.7 %, the total stress C
T
for flows containing passive spheres tends to be slightly lower than that for single-phase flows because the fluid volume fraction is less than unity in particle-laden flows. This trend holds in most cases involving squirmers, except pushers. The scenario for pushers deviates from this pattern at a specified particle volume fraction, exhibiting the smallest u
b
and the highest total stress C
T
, as indicated in
table 2. These findings can be attributed to the self-propelling velocities of the squirmers, which introduces additional kinetic energy into the system and consequently alters the power dynamics of the channel flows and enhances the flow drag. This effect is particularly noteworthy for pushers, as they swim at faster speeds than their counterpart squirmers in the presence of fluid inertia (Chisholm et al. Reference Chisholm, Legendre, Lauga and Khair2016). Notably, a larger C
pI
corresponds to larger values of C
T
and C
pR
, but a smaller value of C
fR
, thereby quantitatively reproducing the analysis depicted in figure 5. It is observed that the fluid Reynolds stress C
fR
for pullers (β = 5) exceeds that for the counterpart pushers (β = −5) at Re
s
= 5. This may be attributed to the magnitudes of vorticity around the body, which serve to strengthen turbulent flows. Specifically, an inertial puller can generate more vorticity than its counterpart pusher (Chisholm et al. Reference Chisholm, Legendre, Lauga and Khair2016). At 𝜑0 = 2.54 %, the results presented in the table provide further quantification of the analysis conducted in the previous section. One of the most significant conclusions confirmed is that pullers demonstrate a drag-reducing effect compared with passive spheres at Re
s
= 5. With an increase in Re
s
, although the total Reynolds stress (C
fR
+ C
pR
) is enhanced, the average velocity of both pullers and fluids decreases.
Profiles of the RMS velocity and Reynolds shear stress for the flow phase (φ
0 = 12.7 %): (a) u
xf
; (b) u
yf
; (c) u
zf
; (d)
$-\langle u_{xf}^{\prime}u_{yf}^{\prime}\rangle$
.

As depicted in figure 7(a–c), our findings corroborate the conclusion that the introduction of passive spheres leads to a decrease in the peak of the streamwise (x) RMS velocity, while simultaneously increasing the wall-normal (y) and spanwise (z) RMS velocities (Shao, Wu & Yu Reference Shao, Wu and Yu2012; Picano et al. Reference Picano, Breugem and Brandt2015). At a volume fraction of 𝜑0 = 12.7 %, most squirmers cause a decrease in the streamwise RMS velocity compared with passive spheres. However, neutral squirmers lead to a slightly higher streamwise RMS velocity than the passive particle case in the vicinity of y + ≈ 42. The mechanism underlying this observation may be attributed to the higher volume fraction of neutral squirmers compared with passive spheres (see figure 4), along with the subtle effect of their self-propelling feature. Throughout the flow profile, the self-propelling mode of squirmers emerges as another critical factor that competes with volume fraction to influence the streamwise RMS velocity. In the near-wall region (e.g. y + ≤ 30), squirmer volume fraction predominates, establishing a negative (positive) correlation between the volume fraction and the streamwise (wall-normal and spanwise) RMS velocity at Re s = 5– higher volume fractions correspond to smaller (larger) velocities. This pattern mirrors what is observed for passive spheres. Conversely, the self-propelling mode prevails in the bulk-flow region (e.g., y + ≥ 120). For instance, despite pushers having a smaller volume fraction than pullers, the streamwise RMS velocity for pushers is smaller than that for pullers. For the puller cases, it is intriguing that an increase in Reynolds number (from 5to 10) results in a decrease in the streamwise RMS velocity but an increase in the wall-normal and spanwise RMS velocities across the entirety of the flow profile. Shao, Wu & Yu (Reference Shao, Wu and Yu2012) suggested that passive spheres can weaken large-scale flow vortices in the streamwise direction and strengthen small-scale vortices when interacting with the wall. Hence, our observation may be attributed to the larger particle inner stress (see figure 5 d) for pullers at Re s = 10, which weakens the large-scale vorticity in the streamwise direction to a greater extent compared with that at Re s = 5. Meanwhile, pullers with larger fluid inertia (e.g. Re s = 10) can induce more small-scale vorticity around the body in all directions than those with small ones (e.g. Re s = 5) (Chisholm et al. Reference Chisholm, Legendre, Lauga and Khair2016). Since the (small-scale) large-scale vorticity dominates the turbulence in the streamwise direction (wall-normal and spanwise directions), the total turbulence intensity is weakened (enhanced). Comparing figures 7(a) and 7(d), the disparities in the fluid Reynolds stresses for different squirmers exhibit notable consistency with those of the streamwise RMS velocities, thus affirming the dominance of turbulence intensity in the streamwise direction, as mentioned above. The RMS velocities and Reynolds stress for the particle phase are shown in figure 8. Its pattern differs from that of the fluid phase. This discrepancy arises because the behaviour of the fluid phase is primarily influenced by the near-wall vortex structure, while particles are closely associated with their own motion (Zhu et al. Reference Zhu, Yu and Shao2018). It is worth noting that at the wall, the RMS velocities do not vanish due to collisions with the wall, and different values for different squirmers indicate varying collision frequencies.
Profiles of the RMS velocity and Reynolds shear stress for the particle phase (φ
0 = 12.7 %): (a) u
xp
; (b) u
yp
; (c) u
zp
; (d)
$-\langle u_{xp}^{\prime}u_{yp}^{\prime}\rangle$
.

Mean direction cosines of the squirmers’ symmetry axes (φ 0 = 12.7 %). (a) Streamwise component (|cos α|). (b) Wall-normal component (|cos β 0|).

4.3. Squirmer orientation, rotation and clustering behaviours
The PDFs of the squirmers’ orientation. (a,c,e) In the bulk-flow region of 135 ≤ y + ≤ 180 (φ 0 = 12.7 %). (b,d,f) In the near-wall region of y + ≤ 45 (φ 0 = 12.7 %). Here cos α, cos β 0 and cos γ denote the direction cosines in streamwise, wall-normal and spanwise components, respectively.

Similar to the analysis of axisymmetric passive spheres, the orientation of a squirmer is described using the direction cosines (|cos α|, |cos β 0|, |cos γ|), where α, β 0 and γ, respectively, denote the angles between the particle symmetry axis and the streamwise, wall-normal and spanwise directions. Figure 9 presents the mean direction cosines of the squirmers’ symmetry axes. In this section, the algorithm of orientation dynamics for nonspherical particles is also employed to handle passive spherical particles (Zhu et al. Reference Zhu, Yu and Shao2018). Initially, the body coordinates of the squirmers and passive spheres are defined, and their orientation is tracked accordingly. It is observed that all average values of the passive spheres are approximately 0.5, indicating the isotropy of the spherical particles and confirming the suitability of this orientation definition for the squirmers. Since each finite-size particle orientation can only be counted once at a specified time (located at its centre), no particle statistics are available near the wall (0 < y + < a + = 22.5). It is observed that the orientation of the squirmers near the wall is not significantly inclined towards the flow direction. As one moves towards the bulk-flow region, pushers and neutrals tend to align their axes more along the flow direction, with pushers showing a more pronounced change. In contrast, pullers tend to favour alignment with the normal direction slightly. Our findings significantly differ from those reported in Zhang et al. (Reference Zhang, Qiu and Zhao2022), where the orientation of microswimmers with an aspect ratio of unity only exhibits minor alterations near the wall and displays isotropy in other regions. This discrepancy may be attributed to the use of a point-particle model, which is suitable for a lower particle volume fraction and does not account for the fluid inertia of the microswimmers and particle–particle interactions. The axisymmetric characteristics of finite-size squirmers and interactions between squirmers and the wall are likely the primary mechanisms contributing to our observed results. The inclusion of self-propelling velocity breaks the geometric isotropy of spherical particles, causing squirmers, especially pushers with lower volume fractions in the bulk-flow region, to adopt specific orientations induced by vorticity in the streamwise direction. Inertial pushers facilitate vorticity convection along their rear (Chisholm et al. Reference Chisholm, Legendre, Lauga and Khair2016; More & Ardekani Reference More and Ardekani2020), enhancing their axial orientation characteristics and making them similar to prolate spheroids. Conversely, inertial pullers engender the accumulation of vortices on both sides of their symmetry axis (Chisholm et al. Reference Chisholm, Legendre, Lauga and Khair2016; More & Ardekani Reference More and Ardekani2020), thereby weakening their axial orientation characteristics and making them similar to oblate spheres. Consequently, the orientation of squirmers in the bulk-flow region aligns with the prevailing notion: particles orient their long axis along the direction of the local fluid vorticity axis (Zhu et al. Reference Zhu, Yu and Shao2018). However, the scenario differs markedly in the near-wall region. Elevated volume fractions of squirmers engender robust hydrodynamic interactions and collisions between particles and with the wall, mitigating the anisotropic properties induced by self-propelling velocity. Near the wall, squirmers tend to exhibit an isotropic orientation distribution akin to that of spherical passive spheres (Zhu et al. Reference Zhu, Yu and Shao2018).
Figures 10(a), 10(c) and 10(e) depict the probability density function (PDF) of the squirmer orientation within the bulk-flow region (135 ≤ y + ≤ 180). The PDF tends towards unity in all directions for passive spheres, further confirming their orientation isotropy. In the bulk-flow region, as shown in figure 10(a), the PDF peaks of pusher orientation occur at cos α = 1 and −1, indicating alignment predominantly parallel to the streamwise direction, consistent with the results shown in figure 9(a). Additionally, the peak at cos α = 1 is higher than that at cos α = −1, suggesting a significantly higher probability of orienting in the positive streamwise direction compared with the negative direction. This orientation may contribute to the reduction in mean flow velocity in the flow direction by pushers (as referenced in figure 3), as their induced flows (pushing for self-propulsion) oppose the streamwise direction. For neutrals, their orientation characteristics in the streamwise direction are less pronounced compared with pushers, showing a monotonic increase across the entire range of −1 ≤ cos α ≤ 1. Interestingly, for the pullers at different Re s , there is a peak at cos α ≈ 0.37 in the streamwise direction, corresponding to an angle of α ≈ 68°. In figures 10(c) and 10(e), within the bulk-flow region, the orientation of squirmers appears somewhat similar in the wall-normal and spanwise directions. However, the notable difference lies in the deviation of the wall-normal cosine peaks for neutrals and pushers from cos β 0 = 0, with deviations of approximately 3° and 5°, respectively. Both neutrals and pushers tend to align perpendicularly to the wall-normal direction, although the pushers show a more pronounced inclination. The cosine orientation values in the wall-normal and spanwise directions for the pullers exhibit minor variation. Two peaks are observed: one parallel to the wall-normal direction (β 0 = 180°) and another at β 0 ≈ 27°. The orientation cosine of the squirmers is symmetric along cos γ = 0, indicating uniformity in fluid alignment in the spanwise direction.
For comparison, figures 10(b), 10(d) and 10(f) depict the PDF of the squirmer orientation within the near-wall region (y + ≤ 45). Unlike pullers or pushers, neutrals do not generate vorticity, making them less likely to be trapped near the wall (Li & Ardekani Reference Li and Ardekani2014). Consequently, the orientation of neutrals does show a pronounced tendency in the streamwise direction. As shown in figure 10(b), there are peaks indicating tendency angles, with α ≈ 35° for pushers and α ≈ 128° for pullers at Re s = 10. In the wall-normal direction (see figure 10 d), pushers and neutrals exhibit peaks at cos β 0 = −1, corresponding to an angle of β 0 = −180°. This indicates that pushers and neutrals tend to swim towards the wall more than pullers, leading to larger aggregations near the wall (see figure 4). Figure 10(f) shows that pushers tend to be perpendicular to the spanwise direction, whereas pullers tend to be parallel to it.
Squirmer mean rotational rates (φ 0 = 12.7 %) in (a) tumbling component and (b) spinning component.

In this section, the rotational behaviour of the squirmer is analysed in terms of ‘spinning’ and ‘tumbling’ actions, with spinning denoting rotation around its symmetry axis (
$\langle w_{z}w_{z}\rangle$
), and tumbling referring to rotation around the other two axes (
$0.5(\langle w_{x}w_{x}\rangle +\langle w_{y}w_{y}\rangle )$
). Figure 11 illustrates the variations in spinning and tumbling speeds across different squirmers, spanning from positions near the wall to the channel centre. Indeed, the rotational speed profiles of spherical passive spheres demonstrate striking similarity, emphasizing their geometric isotropy. The spinning and tumbling velocities of neutral squirmers closely resemble those of passive spheres within the flow field away from the wall, likely attributable to the lack of self-induced vortices in neutral squirmers, similar to passive spheres. Conversely, the vortices generated by pullers and pushers interact with the flow vortices within the channel, leading to pushers displaying increased (decreased) spinning and reduced (enhanced) tumbling speeds. This discrepancy primarily stems from pushers’ inclination to transport their induced vortices downstream, thereby elongating the effective geometric attributes of particles along their symmetry axis, whereas pullers tend to gather vortices around themselves, elongating the effective geometric properties along the other two axes (Chisholm et al. Reference Chisholm, Legendre, Lauga and Khair2016; More & Ardekani Reference More and Ardekani2020). Zhao et al. (Reference Zhao, Challabotla, Andersson and Variano2015) suggested that ellipsoidal particles exhibiting heightened spinning speeds and diminished tumbling speeds can be explained by their tendency to align their long axes with the direction of the channel flow’s vorticity. The facilitation of spinning and tumbling for pushers and pullers, respectively, arises from the alignment of their elongated axes (the symmetry axis for pushers and the transverse axes for pullers) with the direction of the channel flow’s vorticity. As the Reynolds number increases, pullers can induce more vortices around them, leading to enhanced tumbling speeds. Notably, certain squirmers near the wall display minor peaks in rotational speed, likely due to the complex interactions between squirmers and the wall, as well as robust near-field hydrodynamic interactions.
Without considering external flows, previous studies have shown that squirmers can generate large-scale flow structures, thereby inducing aggregation motion (Li & Ardekani Reference Li and Ardekani2014; Lin & Gao Reference Lin and Gao2019; Qi et al. Reference Qi, Westphal, Gompper and Winkler2022; Zhang et al. Reference Zhang, Yu, Ouyang, Xia and Lin2024). Hence, we consider the pair-correlation function g(r), representing the probability of finding a squirmer at a centre-to-centre distance r from another one, to capture the topological structures of squirmer clusters in the turbulent channel flows, and it reads

where r m,n is the distance between squirmers m and n, and V (r m,n ) = (4π/3)[(r m,n + Δr/2)3 − (r m,n − Δr/2)3] is the volume of the spherical shell of radius r m,n and thickness Δr. As shown in figure 12, the pair-correlation functions of squirmers exhibit a first peak around r = 2, reaching a maximum value before rapidly decreasing. This indicates close contact due to pairwise interactions, with the highest peak signifying the most intense interactions between particles. This result correlates with the particle aggregation observed in figure 4(a), where pushers and passive spheres display the highest and lowest peaks near the wall, respectively. A second peak appears around r = 4, consistent with the reported series of double-peak structures at r = 2, 4, 6, 8,… for swimmer clusters (Ishikawa et al. Reference Ishikawa, Locsei and Pedley2008; Lin & Gao Reference Lin and Gao2019; Zhang et al. Reference Zhang, Yu, Ouyang, Xia and Lin2024). Notably, we did not find any double peaks greater than 4, because of our limited channel space. Interestingly, the second peak for the pushers is smaller than that for the passive spheres, in contrast to the situation of the first peak. This may be because, in a limited channel flow, particles with strong pairwise interactions tend to break the formation of larger particle clusters.
Pair-correlation function of squirmers in the turbulent channel flows (φ 0 = 12.7 %).

4.4. Vortex structure
This section investigates the impact of squirmers on vortex structures, aiming to elucidate the potential mechanisms underlying the observed effects of particles on the discussed flow statistics. Figure 13 displays the typical vortex structures in a quarter of the entire calculation domain of the channel flow. We utilize the Q criterion (Q = 1.5) proposed by Hunt, Wray & Moin (Reference Hunt, Wray and Moin1988) for vortex identification, defined as Q = 0.5 (
$\Omega$
ij
$\Omega$
ij
− S
ij
S
ij
), where
$\Omega$
ij
= (u
ij
− u
ji
)/2 and S
ij
= (u
ij
+ u
ji
)/2 are, respectively, the rotation rate tensor and the strain rate tensor. Comparing figures 13(a) and 13(c), the disparity in vorticity is not pronounced. This can be attributed to the fact that neutral squirmers do not inherently generate vortices; rather, the vortices are primarily induced by hydrodynamic interactions and collisions between particles and the wall. Pushers and pullers both generate vorticity themselves. Pushers contribute to the vortex convection opposite to their swimming direction, whereas pullers tend to attract vortices around themselves. As a result, in the bulk-flow region, noticeably higher vorticity induced by pullers compared with pushers can be observed (the abundance of pullers relative to pushers in this region also plays an important role). Meanwhile, a large number of pushers congregate near the wall, giving rise to numerous small-scale vortices (see figure 13
e). These small vortices are generated both by pushers and by the weakening and fragmentation of large-scale vortices, resulting in a minimal flow RMS velocity in the near-wall region compared with that for other squirmers (see figure 8
a). The orientation and distribution characteristics of the particle phase are depicted in figure 13(b,d,f,h). Upon comparison of figures 13(d), 13(f) and 13(h), it is confirmed that the vorticity in the central region of the flow field is primarily induced by the squirmers themselves: the vorticity generated by neutrals is minimal, whereas that generated by pushers is largely concentrated at their tails, accompanied by no discernible pattern (appearing uniform) in the distribution of vorticity around pullers.
Vortex structures of the particle-laden channel flows (φ 0 = 12.7 %) filled with (a,b) passive spheres, (c,d) neutral squirmers, (e,f) pushers and (g,h) pullers. (a,c,e,g) Oblique view with removing particles. (b,d,f,h) Side view in the spanwise direction with particles. The red caps on the squirmers indicate their heads (orientation). A quarter of the calculation domain is displayed to enhance clarity when observing the vortex structure and particle orientation. Note that the colour of the vortices represents the fluid streamwise velocity, and darker means a larger streamwise velocity.

5. Conclusion
We have numerically investigated the modulation and transport characteristics of finite-size microorganisms in turbulent channel flows using a DF-FD method incorporating the squirmer model. The study systematically examines the effects of suspended squirmers on the fluid’s mean and RMS velocities, vortex structures and the properties of the squirmer velocities, concentration distribution, orientation and rotation rate. This analysis is conducted at a friction Reynolds number of 180, considering two squirmer volume fractions (12.7 % and 2.54 %) and a blocking ratio of 0.125.
Our results show that, at a high volume fraction (𝜑0 = 12.7 %), pushers induce a more pronounced decrease in the velocity profile compared with neutral squirmers and pullers. This disparity arises from the distribution of squirmers near the wall, where a higher density of particles leads to slower velocity profiles. Generally, squirmers, particularly at higher volume fractions, weaken the fluid velocity profile more significantly than passive particles. This phenomenon primarily arises due to their pronounced tendency to aggregate near walls, forming a dense particle layer. Such wall-attraction behaviour of microswimmers is notable in practical contexts, as it likely contributes to the blockage of sewage pipelines, where volume fractions of planktonic microorganisms are frequently present. At a low volume fraction (𝜑0 = 2.54 %), similar but reduced effects on the velocity profiles are observed, indicating less impeding by the squirmers compared with the high volume fraction. Notably, at this lower concentration, pullers slightly enhance the velocity profile across the channel compared with passive spheres.
Regarding the stress of the squirmer-laden turbulent channel flows, different self-propelling modes affect the particle inner stress τ pI in the near-wall regions differently, especially at the high particle volume fraction (𝜑0 = 12.7 %). Pushers contribute to the highest peak of particle inner stress τ pI compared with other squirmers. This pattern may be attributed to the combined effects of particle accumulation behaviours (the distribution of volume fraction) and the self-propelling features. Significantly, the increase in τ pI is primarily at the expense of diminishing the fluid Reynolds stress τ fR . This provides valuable insights into the hydrodynamics of mixed systems containing planktonic microbial particles. At 𝜑0 = 2.54 % (Re s = 5), the peak of particle inner stress τ pI for the puller case is smaller than that of passive spheres, resulting in a greater fluid Reynolds stress τ fR . This observation could be a key factor contributing to the higher flow profile velocity observed for pullers compared with passive spheres.
Regarding the squirmer dynamics, the orientation of the squirmers near the wall does not significantly incline towards the flow direction, in contrast to the rheotaxis observed in microorganisms driven by shear flow in laminar conditions near the wall (Zaferani, Cheong & Abbaspourrad Reference Zaferani, Cheong and Abbaspourrad2018; Wheeler et al. Reference Wheeler, Secchi, Rusconi and Stocker2019; Omori et al. 2020). In the bulk-flow region, pushers and neutrals align their axes more along the flow direction, with pushers showing a particularly strong alignment. In contrast, pullers tend to favour alignment with the normal direction slightly. The spinning and tumbling velocities of neutrals closely resemble those of passive spheres except near the wall, likely attributable to the lack of self-induced vortices in neutrals, similar to passive spheres. In contrast, the vortices generated by pullers and pushers interact with the flow vortices within the channel. As a result, pushers exhibit increased spinning and reduced tumbling speeds, while pullers show decreased spinning and enhanced tumbling speeds. Notably, some squirmers near the wall display minor peaks in rotational speed, likely due to complex colliding between squirmers and the wall, as well as strong near-field hydrodynamic interactions.
The current study has focused solely on two kinds of volume fractions for squirmers, primarily due to computational constraints and limited resources. Further investigations across a broader parameter range are necessary to comprehensively understand the interactions between finite-size squirmers and turbulent channel flows.
Funding
The authors would like to thank the National Natural Science Foundation of China (grant nos. 12302333; 12132015; 12372264; 12172327), Zhejiang Provincial Natural Science Foundation of China (grant no. LQ24A020003) and International Sci-tech Cooperation Projects under the ‘Innovation Yongjiang 2035’ (grant no. 2024H016).
Declaration of interests
The authors report no conflict of interest.
Data availability statement
The data that support the findings of this study are available from the corresponding author upon reasonable request.





